Journal of Experimental Biology · 2004 · 176 citations · 66 references
EngineeringEntomologyMechanical EngineeringAerodynamicsFlight DynamicsFlight MechanicsStroke MechanicsWing AerodynamicsAerodynamic ModelingElectromechanical ModelingFlapping FlightFlapping Wing AerodynamicsFlapping Wing LiftWake InterferenceWing Beat KinematicsWing Wake Dynamics
Insects with two pairs of wings must contend with forewing wake interference over the hindwing, and the phase relationship between fore‑ and hindwing strokes can influence total lift, but its effect is hard to assess due to complex wake patterns. The study investigates how altering the fore‑ and hindwing stroke‑phase relationship during hovering flight affects the aerodynamic performance of each wing using a dynamically scaled electromechanical insect model. The authors employ a dynamically scaled electromechanical insect model to systematically vary the fore‑ and hindwing stroke‑phase relationship and measure the resulting aerodynamic performance. Varying the fore‑ and hindwing phase difference leaves forewing lift unchanged while hindwing lift can change by up to a factor of two, with optimal performance occurring when the hindwing leads the forewing by about a quarter cycle, a relationship that matches natural insect phase shifts and suggests neuromuscular control of phase can modulate ipsilateral lift without altering wingbeat kinematics.
SUMMARY Insects flying with two pairs of wings must contend with the forewing wake passing over the beating hindwing. Some four-winged insects, such as dragonflies, move each wing independently and therefore may alter the relative timing between the fore- and hindwing stroke cycles. The significance of modifying the phase relationship between fore- and hindwing stroke kinematics on total lift production is difficult to assess in the flying animal because the effect of wing-wake interference critically depends on the complex wake pattern produced by the two beating wings. Here we investigate the effect of changing the fore- and hindwing stroke-phase relationship during hovering flight conditions on the aerodynamic performance of each flapping wing by using a dynamically scaled electromechanical insect model. By varying the relative phase difference between fore- and hindwing stroke cycles we found that the performance of the forewing remains approximately constant, while hindwing lift production may vary by a factor of two. Hindwing lift modulation appears to be due to two different fluid dynamic phenomenons: leading edge vortex destruction and changes in strength and orientation of the local flow vector. Unexpectedly, the hindwing regains aerodynamic performance near to that of the wing free from forewing wake interference, when the motion of the hindwing leads the forewing by around a quarter of the stroke cycle. This kinematic relationship between hind- and forewing closely matches the phase-shift commonly used by locusts and some dragonflies in climbing and forward flight. The experiments support previous assumptions that active neuromuscular control of fore- and hindwing stroke phase might enable dragonflies and other functionally four-winged insects to manipulate ipsilateral flight force production without further changes in wing beat kinematics.
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Wing Rotation and the Aerodynamic Basis of Insect Flight
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